NATURAL RESOURCES CONSERVATION SERVICE Plant Materials Technical Note NV-58 November 2015

Milkweed Pollination Biology (Asclepias spp.)

Eric P. Eldredge, Manager, Great Basin Plant Materials Center, Fallon, Nevada

Monarch butterfly (Danaus plexippus) conservation depends on conservation of the milkweeds that are the host plant for monarch larvae. Across much of its range in the western U.S., showy milkweed (Asclepias speciosa Torr.) is an important source of nectar for adult monarch butterflies and is the preferred host plant for monarch larvae (Borders, et al., 2012). Milkweed flowers bloom in umbels (Figure 1), which are clusters of individual flowers on stems that emerge from a common point (Young-Mathews and Eldredge, 2012).

Figure 1. An umbel of showy milkweed flowers in full bloom.

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Each flower has a calyx, corolla, and corona, which surround a central gynostegium. The gynostegium is formed from the fused, highly modified male (anthers and filaments) and female (stigmas and styles) floral parts. At the top of the gynostegium is a style-head, formed by fusion of the apices of two styles. The corolla is the outer and lower part of the flower and resembles petals (Figure 2). In some milkweed species the corolla is bent backwards (reflexed).

A

B

D C

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Figure 2. A top view of a showy milkweed flower showing the corolla (A), and the corona with five hoods (B), each with a prominent horn (C). A dark brown corpusculum (D) is visible at the top of three of the stigmatic slits (E). The corona is the showy, upper part of the flower. Five hoods surround the corona and tempting pools of luscious nectar form at the base of each hood. In some species, each hood has a prominent horn. Together these slick, waxy floral structures manipulate the behavior of insects to achieve pollination.

Flowers of different species of milkweed differ in size, color, and fragrance, but all produce their pollen in waxy sacs called pollinia. The pollinia are located in two anther pouches adjacent to vertical stigmatic slits of the flowers (Figure 3). Pairs of adjacent pollinia are connected to each other by translator arms from a clamp located in the middle, called the corpusculum (Bookman, 1981). The complete structure is called a pollinarium (Figure 4). A very similar pollinarium structure also evolved in a monocotyledonous group of insect pollinated plants: the orchids. Because of their unusual structure and relatively large size, the flowers of showy milkweed require relatively large insects for effective pollination. 2

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Figure 3. A magnified view of a dissected showy milkweed flower, with the anther pouch walls (AA) removed to show a pollinarium and the position of the pollinia (BB), translator arms (CC), and the corpusculum (D) at the top of the stigmatic slit (E) formed by the guide rails (FF). Insects that visit a flower to drink nectar struggle to grasp the slippery surfaces and may accidently slip their leg, tarsus, mouthpart, or other appendage into the opening at the bottom of the stigmatic slit. This slit is formed by guide rails, which are lined with bristles that prevent the insect moving its appendage any direction but up. The top of the slit leads into the opening of the corpusculum, which has hard, sharp inside edges that taper together at the top. The corpusculum clamps firmly to the insect by pinching onto the insect's appendage. In its struggles to escape, if the insect is large enough, it can withdraw the paired pollinia from the anther pouches and fly away (Figure 5).

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Figure 4. A pollinarium of showy milkweed on the edge of a millimeter scale, and an exposed pollinarium in the background.

Figure 5. An unidentified large wasp nectaring on narrow leaf milkweed, bearing many pollinaria attached to the hairs on its legs (at arrows). 4

During flight, the translator arms dry and bend to change the orientation of the pollinia (Bookman, 1981). When the insect visits another milkweed flower, a pollinium may slip into one of the stigmatic slits and be captured inside the stigmatic chamber. The insect must then break the translator arm to escape, leaving the pollinium in the stigmatic chamber where the pollen grains can germinate to fertilize the flower. Deep inside the gynostegium, two future seedpods (follicles) have already developed the unfertilized ovules by the time the flower blooms. One follicle can be pollinated by a pollinium inserted into any of three adjacent stigmatic slits, and the other follicle is connected to the two remaining stigmatic chambers (Sage et al., 1990). After pollination occurs, the other flowers on the umbel wither and fall off. Not all insects that visit to showy milkweed to drink nectar are large enough, strong enough, and clumsy enough to pollinate it effectively. Introduced European honeybees can pollinate showy milkweed effectively because they are so numerous. Showy milkweed provides abundant nectar for honeybees over a long bloom period. However, due to their small size relative to a showy milkweed flower, occasionally a honeybee may be trapped on the flower (Figures 6 and 7).

Figure 6. Some bees and smaller insects, unable to withdraw the pollinarium, may lose a limb, or be trapped on the showy milkweed umbel, like this honeybee.

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Figure 7. The headless carcass of a trapped honeybee, one leg lost and still not free. A catena of corpuscula is visible to the right of her trapped tarsus (at arrow). After insertion of a pollinium into a stigmatic slit, if the insect is strong enough to break the translator arm and escape, the broken stub is the perfect size, shape, and orientation to catch another corpusculum. This may repeat until chains (catena) of corpuscula are formed, in a process called concatenation (Peter and Shuttleworth, 2014). More research is needed to determine if population declines in the insect species that are the most effective pollinators of milkweed (Wiemer et al., 2012) have contributed to the nationwide decline in milkweed, and consequent decline in monarch butterfly habitat. The next time you see a milkweed seedpod (follicle) think about the hapless insect who, in exchange for a rich reward of carbohydrate, removed a pollinarium and flew to a flower on another milkweed, where a pollinium chanced to slip into the stigmatic slit. Without a sufficient diversity of pollinator species to accomplish this peculiar and elaborate pollination process, the showy milkweed populations that support the monarch butterfly could continue to decline. Showy milkweed grows best in well-drained soil in full or nearly full sun, in pastures, meadows, forest clearings, untilled fields, roadsides, and ditch banks, from sea level to 6,250 ft. (Stevens and Anderson, 2005). Seed is routinely commercially available. Whenever possible, showy milkweed should be considered as a component in pollinator habitat plantings, such as backyard gardens, along field borders, and edges of hedgerows. Showy milkweed should be considered for roadside beautification projects when it can be planted some distance away from the flow of traffic. The flowers bloom from May through September, providing nectar to a diverse community of pollinating insects.

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Literature Cited

Bookman, S.S. 1981. The floral morphology of Asclepias speciosa (Asclepiadaceae) in relation to pollination and a clarification in terminology for the genus. American Journal of Botany 68:675-679. Borders, B., E. Eldredge, E. Mader, and C. Burns. 2012. Great Basin Pollinator Plants: Native Milkweeds (Asclepias spp.) The Xerces Society for Invertebrate Conservation, Portland OR, in collaboration with USDA-NRCS Great Basin Plant Materials Center, Fallon, Nevada. NVPMC Technical Note No. 56. Peter, C.I., and A. Shuttleworth. 2014. Catching on to concatenation: evidence for pre-pollination intra- sexual selection in plants. New Phytologist 203:4-6. Sage T.L., S.B. Broyles, and R. Wyatt. 1990. The relationship between the five stigmatic chambers and two ovaries of milkweed (Asclepias amplexicaulis Sm.) flowers: a three- dimensional assessment. Israel Journal of Botany. 39:187-196. Stevens, M., and M.K. Anderson. 2005. Plant Guide for showy milkweed. USDA NRCS National Plant Data Center. Revised May 2006. 4 pp. Wiemer, A.P., A.N. Sersic, S. Merino, A.O. Simoes, and A.A. Cocucci. 2012. Functional morphology and wasp pollination of two South American asclepiads. Annals of Botany 109(1) 77-93. Young-Mathews, A., and E. Eldredge. 2012. Plant Fact Sheet for showy milkweed. USDA NRCS Corvallis Plant Materials Center and Great Basin Plant Materials Center. Corvallis, OR and Fallon, NV. August 2012. 2 pp. Citation Eldredge, E. 2015. Milkweed Pollination Biology. USDA NRCS Technical Note NV-58, Great Basin Plant Materials Center, Fallon, NV. The U.S. Department of Agriculture (USDA) prohibits discrimination against its customers, employees, and applicants for employment on the bases of race, color, national origin, age, disability, sex, gender identity, religion, reprisal, and where applicable, political beliefs, marital status, familial or parental status, sexual orientation, or all or part of an individual's income is derived from any public assistance program, or protected genetic information in employment or in any program or activity conducted or funded by the Department. (Not all prohibited bases will apply to all programs and/or employment activities.) If you wish to file an employment complaint, you must contact your agency's EEO Counselor (PDF) within 45 days of the date of the alleged discriminatory act, event, or in the case of a personnel action. Additional information can be found at http://www.ascr.usda.gov/complaint_filing_file.html. If you wish to file a Civil Rights program complaint of discrimination, complete the USDA Program Discrimination Complaint Form (PDF), found at http://www.ascr.usda.gov/complaint_filing_cust.html, or at any USDA office, or call (866) 632-9992 to request the form. You may also write a letter containing all of the information requested in the form. Send your completed complaint form or letter to us by mail at U.S. Department of Agriculture, Director, Office of Adjudication, 1400 Independence Avenue, S.W., Washington, D.C. 20250-9410, by fax (202) 690-7442 or email at [email protected]. Individuals who are deaf, hard of hearing or have speech disabilities and you wish to file either an EEO or program complaint please contact USDA through the Federal Relay Service at (800) 877-8339 or (800) 845-6136 (in Spanish). Persons with disabilities, who wish to file a program complaint, please see information above on how to contact us by mail directly or by email. If you require alternative means of communication for program information (e.g., Braille, large print, audiotape, etc.) please contact USDA's TARGET Center at (202) 720-2600 (voice and TDD). For any other information dealing with Supplemental Nutrition Assistance Program (SNAP) issues, persons should either contact the USDA SNAP Hotline Number at (800) 221-5689, which is also in Spanish or call the State Information/Hotline Numbers. For any other information not pertaining to civil rights, please refer to the listing of the USDA Agencies and Offices for specific agency information.

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